The overall goal of the proposed research is a quantitative description of the structural and physiological constraints on low-frequency selectivity in the vertebrate auditory system. In particular, the primary objectives of the proposed research are to gain an understanding and appreciation of the mechanical and electrical factors underlying airborne, substrate-borne and combination (bimodal) stimulus reception, and to provide further insight into the mechanisms underlying stimulus interactions which affect tuning in the vertebrate inner ear. To accomplish these objectives, a series of five detailed investigations will be performed in order to a (a) directly measure the motion of the middle ear ossicles in amphibians in response to airborne sound, substrate-borne vibration and bimodal stimulation, and thus more precisely define the role of these structures in low-frequency reception, (b) characterize the airborne and seismic response properties of the middle ear ossicles of two other "low- frequency" animals- the common and golden mole- and thus extend our observations to fossorial mammals, (c) quantify the extent to which the tectorial membrane responds to low-frequency sound and vibration in order to elucidate the role of this structure in bimodal processing, (d) systematically compare synaptic release in low-frequency (bimodal) and high-frequency (unimodal) hair cells from the amphibian papilla by tracking correlated capacitance changes in response to depolarization, and (e) extend our investigation of the nonlinear interactions between acoustic and seismic stimuli to the bimodal fibers in the eighth nerve. The data that result from this integrative structure-functional and neuroethological approach will be rich in implications regarding the anatomical and neural substrates underlying the processing of sound-vibration complexes; thus this work is expected to provide a framework for understanding the relationship between air-conducted and bone-conducted sound transmission in animals, including humans.